Evolution of Altruism

In evolutionary biology, altruism has a special meaning.
As well as benefiting others, altruistic behaviour must incur a
definite loss to the individual carrying it out.
It is usually, but not exclusively observed within members
of the same species. For example,
a bee which stings an agressor gives up its life,
but helps the colony survive by doing so.
Altruism
so puzzled biologists schooled on a competitive interpretation of evolution
that E.O. Wilson, echoing the sentiments of many called it "the culmination mystery of all biology".

The first biologist to advance an explanation of altruism was William Hamilton.
In 1963 he showed how kin selection (in which family members behave altruistically towards one another) could evolve.
Robert Trivers showed in 1971 how
repeated altruistic exchanges
could evolve between unrelated members of the same species, or even of different species.
Subsequently more complicated theories have arisen, including
multi-level selection which posits simultaneous selection pressure
on different levels, and
costly signalling theory which suggests that altruism establishes reputation.
It seems a fair conclusion that
altruism has evolved through a mixture of all of these mechanisms,
as well as more subtle ones as yet undiscovered.

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